Project Summary
Intellectual merit: Ecologists have long been challenged to understand species distributions and the
ecological factors that maintain species range limits. Montane regions have allowed ecologists to study
range limits for multiple species simultaneously because habitats and species composition often change
abruptly along altitudinal gradients. Tropical montane regions should be key landscapes in this area of
research because of their high species diversity and the concentration of range limits at small spatial
scales along altitudinal gradients, which offers unprecedented opportunity to study replicated patterns.
Also, though not well explored, tropical montane regions have revealed several distinctive patterns of
species association, contrary to the long championed individualistic Gleasonian patterns of temperate
communities. My dissertation research focuses on two such patterns found in birds: (1) replacements in
closely related species along altitudinal gradients and (2) correspondence of bird communities to plant
community composition. Because it is difficult to examine mechanisms that limit species distributions
when looking at the community level for birds and plants, I have chosen to focus on a particular subset of
these communities for which we might expect strong associations along gradients: hummingbirds and
flowering plants. Hummingbirds are primarily nectar-feeding birds that serve as important pollinators for
flowering plants. In this group, studies have shown many cases of highly specialized plant-pollinator
relationships. The highest diversity of hummingbirds is found in the forests of the Andes Mountains in
South America, where species often have altitudinal distributions of a few hundred meters. Despite their
well-studied specialization to flowering-plants, no one has examined how relationships between
hummingbirds and flowering plants may constrain hummingbird species' altitudinal distributions. One
enormous advantage of hummingbirds is that they leave a record of which flowers they have visited
through the accumulation of pollen on their bills and heads, thus making costly studies that follow
hummingbirds as they move through the environment unnecessary. In the proposed project, more than
400 pollen samples already collected from bills of captured hummingbirds along an altitudinal gradient in
the Manu Biosphere Reserve of Peru will be identified to compile a list of flowering plants visited by each
of 38 species of hummingbirds. These data, combined with data on the occurrence of those hummingbird
and flowering-plant species along the same gradient, will be used to address the following hypotheses:
(1) range limits in hummingbirds specialized to few flowering-plant species coincide with limits in the
distribution of their flowering plants; (2) hummingbirds with broad altitudinal distributions change their use
of flowering-plant species with altitude, making them specialists on different flowering-plant species within
parts of their range; and (3) hummingbirds specialized to fewer flowering-plant species are constrained to
narrower altitudinal ranges compared to generalist hummingbirds. This will be the first study to examine
how ranges of one taxonomic group (plants) affect the distribution of another taxon (birds).
Broader impacts: Tropical mountains have been targeted as regions where climate change will have
high impacts on flora and fauna. This analysis of flowering-plant specialization and range limits in
hummingbirds could reveal potential limitations on this diverse family in their ability to respond to
changing temperature regimes. The proposed work represents an ongoing collaboration with the
Neotropical Paleoecology Laboratory, directed by Dr. Mark Bush at the Florida Institute of Technology,
whose focus has been to develop a reference collection and pollen database for Neotropical flowering
plants, with a strong concentration on Andean flora of southeastern Peru. This collaboration has fostered
field research experience for an undergraduate student at Florida Tech who worked with me during the
collection of pollen samples in 2006. The proposed research will create another undergraduate research
position in this lab facility to process the pollen samples and will include a trained palynologist from
Cuzco, Peru, to identify the pollen. Any pollen samples new to the Neotropical pollen collection will
contribute to a growing online database. Many undergraduate students have already participated in data
collection and analysis during the field component of this project from 2005-2008, including eight female
Peruvian biology students, one female Colombian undergraduate, as well as two undergraduates and one
high school student from the United States. Three of the Peruvian students are completing their
"licenciatura" projects using data from this project, under the mentorship of the Co-PI. The proposed
research, through additional collaboration with botanists from Dr. Miles Silman's Lab in Wake Forest
University, will be used to produce a hummingbird-flowering plant photo guide, featuring hummingbird
species for which pollen is being analyzed in this study along with their primary flowering plants in Manu.
Another product resulting from this project will be a set of three compact discs of recorded bird species
from the montane forests of Manu, which are currently being organized by an undergraduate student.
The hummingbird-flowering plant photo guide and compact discs will be invaluable learning resources for
researchers beginning investigation with birds of the Peruvian Andes.

Project Description (Including Results from Prior
NSF Support) (not to exceed 15 pages) (Exceed only if allowed by a
specific program announcement/solicitation or if approved in
advance by the appropriate NSF Assistant Director or designee)

References Cited

Biographical Sketches (Not to exceed 2 pages each)

Budget
(Plus up to 3 pages of budget justification)

Current and Pending Support

Facilities, Equipment and Other Resources

Special Information/Supplementary Documentation

Appendix (List below. )
(Include only if allowed by a specific program announcement/
solicitation or if approved in advance by the appropriate NSF
Assistant Director or designee)

Appendix Items:

*Proposers may select any numbering mechanism for the proposal. The entire proposal however, must be paginated.
Complete both columns only if the proposal is numbered consecutively.

J.E. Jankowski

USING HUMMINGBIRDS TO TEST THE LINK BETWEEN BIRD AND PLANT
DISTRIBUTIONS IN TROPICAL MONTANE FOREST

Ecologists have long struggled to understand species distributions and the ecological factors
that maintain species range limits (Brown et al. 1996, Case and Taper 2000, Case et al. 2005,
Holt and Keitt 2005). An underlying problem is that range limits of different species do not co-
occur in space, making it logistically difficult to study mechanisms for more than a few species
at a time. Montane regions provide a welcome exception because habitats and species
composition often change abruptly along altitudinal gradients, giving ecologists the opportunity
to study distributions of many species simultaneously. For example, Whittaker (1967) used tree
distributions along temperate altitudinal gradients to test the classic "individualistic" and
"community" hypotheses of H.A. Gleason and F.E. Clements, respectively, which described how
species are distributed with respect to each other along gradients.
Because of their high species diversity and high density of range limits at small spatial
scales, tropical mountain ranges have enormous, mostly untapped potential as study systems
for understanding patterns of species distributions. In birds, hundreds of species can inhabit a
single tropical mountainside, with many species having altitudinal ranges of only a few hundred
meters (Stotz et al. 1996). Curiously, few studies have focused on tropical montane regions to
address questions of species distributions and range limits (Terborgh 1971, Hernandez et al.
2008, Buermann et al. 2008).
Besides offering a greater amount of "raw material" of diversity to describe replicated
patterns in species distributions, tropical montane communities have often revealed a very
different pattern than temperate montane communities. While studies in temperate montane
regions have largely concluded that species in a community are distributed individualistically
with altitude, as originally proposed by Gleason (Gleason 1939, Whittaker 1956, Peet 1981),
studies of montane species in the tropics reveal many situations in which multiple species'
range limits co-occur along altitudinal gradients (but see Terborgh 1971, Navarro 1992,
Patterson et al. 1998). One such case common in birds is that of "species replacement", in
which closely related species show adjacent non-overlapping distributions. The classic
interpretation of this pattern is that interspecific competition underlies replacements by
preventing the coexistence of strong competitors (Terborgh and Weske 1975), although
community-wide patterns suggesting competition have rarely been critically assessed or verified
experimentally (but see Robinson and Terborgh 1995, Martin and Martin 2001). Alternatively,
multiple species may show associated distributions because they are responding to sudden
changes in habitat structure or plant communities along the gradient (e.g., ecotones; Terborgh
1985, Navarro 1992, Jankowski etal. in press). In these cases, species should have abrupt
range limits that coincide with sharp transitional zones in habitat structure or in plant
composition. A third explanation is based on specialized plant-animal interactions, which are
prominent features of tropical forests (e.g., plant-pollinator relationships; Janzen 1975,
Dziedzioch et al. 2003). We might expect specialized pollinators, for example, to have
altitudinal distributions tightly associated with the distributions of their host plants (Stiles 1975).

For my dissertation, I am documenting patterns of species distributions for a tropical
montane bird community in southeastern Peru and investigating several processes that could
influence species range limits along a 2600-m altitudinal gradient. I have focused on two
hypotheses:
(1) Replacements in closely related species along altitudinal gradients are maintained by
competitive interactions between species. For territorial animals such as birds, interspecific
competition is often expressed as interspecific territoriality (i.e., territorial defense against
individuals of other species). I have used heterospecific playbacks of songs conducted within

J.E. Jankowski

territories of species showing altitudinal replacements and measured behavioral responses to
test for the presence of interspecific territoriality in these species.
My data support this hypothesis: species respond aggressively to song playbacks of closely
related species (i.e., congeners). In many cases, this response is nearly as strong as the
individuals' response to conspecific songs (i.e. songs of their own species). Interestingly, the
strength of behavioral response of an individual to a congener song playback decreases as one
moves away from the zone of replacement along the gradient, suggesting that the development
of such behavioral responses requires contact with individuals of the other species (Figure 1).

Figure 1. Closest approach to speaker (an aggressive response to song playbacks) in
Henicorhina leucophrys for an observation period (no stimulus or control) and congener and
conspecific song playbacks. The congener song playback used H. leucosticta. Bars show
categories of distance of the focal territory to the nearest congener territory. When territories
are close to the replacement zone, H. leucophrys responds very strongly to congener songs,
but the strength of response declines with increasing distance from the replacement zone
(GLM: F = 75.8, df = 6, p<0.0001; the following variables were significant at the p<0.02 level:
trial (i.e., observation, congener or conspecific), distance from nearest congener, and
interaction between trial and distance).

(2) Bird and plant communities show similar peaks of species turnover, or shared zones of
rapid transition, along altitudinal gradients and show high correspondence in species
composition. I am using avian censuses that I conducted along this altitudinal gradient, and
collaborating with botanists working in the same region, to determine the degree to which
elevational changes in plant communities match (and presumably drive) elevational limits of
birds. These comparisons include examining patterns in species turnover along the gradient
and the community-wide correspondence of these taxa in this landscape e.g., are sites that
are more similar in bird composition also more similar in plant composition? Data to test this
hypothesis are currently being analyzed.
These hypotheses are not mutually exclusive because species replacements mediated by
interspecific competitive interactions could occur at boundaries between different habitats (e.g.,
ecotones) along gradients.
The proposed research will focus on a particular subset of birds and plants for which
we might expect strong associations between bird and plant distributions along
gradients: hummingbirds and flowering plants. In Neotropical birds, hummingbirds are
important pollinators for flowering plants and have often developed specialized mutualistic

J.E. Jankowski

relationships with plant species. It is possible that range limits in this avian foraging guild (i.e.,
nectarivores) are resource driven, such that distributional limits of specialists should coincide
with those of their flowering-plant food resources along the gradient. If any group should show a
correspondence between plant and bird distributions, it should be this group. If there are no
associations between plants and birds in this group, then there might be little chance of finding
these associations in any other group.

Hummingbirds and flowering-plants as a system for examining plant-based
distributional limits along gradients

Interactions between birds and plants have been a natural starting point for biologists aiming
to understand habitat associations and species distributions because plants provide food
resources as well as habitat structure used for foraging and shelter for many species (Cody
1985, Estades 1997, Lee and Rotenberry 2005). Most bird-plant interactions are generalized,
and though the community composition of plants and birds may show an overall
correspondence, it would be unlikely that the distribution of a given species of bird would closely
match that of any single plant species. However, there are specialized relationships that have
developed between birds and plants, especially in plant-pollinator systems (e.g., hummingbirds
and flowering plants). Such plant-pollinator relationships offer promising avenues to investigate
the influence of resource specialization on altitudinal distributions.
Hummingbirds (Family Trochilidae) make up the second largest family of birds in the New
World, totaling 328 species, the large majority of which are tropical and montane. They are
highly evolved nectarivores, dependent upon carbohydrate-rich nectar of flowering plants for
90% of their energetic requirements, and they serve as important pollinators for a variety of
flowering-plant species (Schuchmann 1999). Consequently, flowering plants have evolved
colorful flowers to attract their avian mutualists. Likewise, morphological traits of hummingbirds
(e.g., bill size and shape) closely match flower morphology, often making them specialized to a
limited spectrum of available nectar resources in the environment. Although the hummingbird-
flowering plant linkage represents one of the strongest of bird-plant relationships, the degree of
specialization of hummingbirds to flowering plants varies remarkably among species, ranging
from generalists that feed almost non-selectively on flowers, to coevolved specialists that utilize
only a few species of flowering plants, which precisely match the hummingbird's bill and body
morphology (e.g., Ensifera ensifera, the Sword-billed Hummingbird, and its primary nectar
source, Passiflora mixta, Lindberg and Oleson 2001; Stiles 1975, Stiles 1985). At the
community level, studies of tropical hummingbird pollinator-plant relationships and specialization
(mostly from Central America) have classified hummingbirds based on bill and flower
morphology (Stiles 1985) or using hummingbird foraging behavior (e.g., traplining hummingbirds
versus territorialists; Feinsinger and Colwell 1978). These and other studies report high levels
of plant-pollinator coevolution within multiple hummingbird communities in tropical lowland and
montane forest. However, no study has examined how this specialization on flowering-plant
resources may constrain species distributions in hummingbirds.
The proposed research will determine whether species distributions and range limits in
hummingbirds are associated with their flowering-plant resources along an altitudinal gradient in
southeastern Peru. Data for this study, collected during my field research along an altitudinal
gradient in the Manu Biosphere Reserve, offer a promising new area of investigation of the
factors that influence species range limits in birds. Additionally, this research takes a novel
approach that overcomes previous difficulties in examining resource specialization in
hummingbirds. In particular, prior studies have used observations of hummingbirds at flowering
plants in their environments to describe plant-pollinator relationships (e.g., Dziedzioch et al.
2003). This approach is problematic, especially in diverse tropical systems, because it requires
repeated observations of all species of flowering plants in a region. There is a simple solution-

J.E. Jankowski

because hummingbirds carry, on their bills and foreheads, pollen of the flowering plants they
visit, one can sample and identify the pollen from captured individuals to determine which
flowers are being visited. With this in mind, I collected pollen samples from the bills of captured
hummingbirds during mist-netting surveys along the altitudinal gradient. By collaborating with
specialists that can identify the pollen grains collected from each individual, I can create a list of
flowering-plant species used by each species of hummingbird. These data, combined with
distributional information from censuses for the hummingbirds and flowering plants along the
gradient will be used to address the following hypotheses:

1) Range limits in hummingbirds that are specialized to few flowering-plant species coincide
with limits in the distribution of their flowering plants. For all of the species sampled along
the gradient, I will test whether, and how often, the altitudinal range limits of hummingbirds
and the range limits of their primary flowering plants coincide, as determined from our
censuses of birds and plants in the region.
2) For hummingbird species with broad altitudinal distributions, hummingbirds change their use
of flowering-plant species with altitude, making them specialists on different sets of
flowering-plant species within parts of their altitudinal range. The capture of each
hummingbird during mist-netting can be pinpointed to a specific location along the gradient,
so that the use of different flowering plants by individuals of each hummingbird species can
be mapped with elevation.
3) Species that are more specialized, or that use fewer species of flowering plants, are
constrained to narrower altitudinal ranges compared to generalist hummingbirds. Using data
on the number of flowering-plant species utilized and the altitudinal distributions of
hummingbirds, I will test whether resource specialization is associated with restricted
species distributions.

Data Collection and Analyses

Mist-netting surveys and collection of pollen along the gradient-Hummingbird pollen was
collected during mist-netting surveys along the altitudinal gradient in Manu Biosphere Reserve
between 850 3400m between the months of August and November, 2006 2007, falling
within the avian montane breeding season. During this time, 42 groups of mist-nets, with ten
nets per group, were run for three days each, yielding a total of 9,360 net hours. All netting
sites were mapped using a GPS, allowing us to pinpoint the capture location and elevation for
each individual to within 7 meters. At least one netting group was placed every 200m altitude
along the gradient. Mist-netting provided almost 1,000 captures of 39 species of hummingbirds
and 38 species for which pollen and morphological data could be collected and recorded (Table
1). These species are distributed rather evenly among foothill, middle elevation, and highland
sites. Pollen was collected from captured hummingbirds using a slice of slightly adhesive
gelatin, which was held with tweezers and rubbed over the entire surface of the bill, forehead
and chin. After transferring pollen from the hummingbird to the gelatin, the gelatin was placed
on a glass microscope slide, heated with a small flame until the gelatin liquidized, and covered
with a glass cover slip to protect the pollen sample from contamination.

Pollen identification-Hummingbird pollen samples will be identified between June -
September of 2009 within the Neotropical Paleoecology Laboratory, directed by Dr. Mark Bush
at the Florida Institute of Technology, where resources and equipment will allow the most
efficient and complete identification of Andean pollen. In this laboratory there is a large
reference collection of pollen from the Southeastern Peruvian Amazon and Andes and a
Neotropical pollen database for more than 1,000 species. The pollen samples will be processed
and remounted onto microscope slides for identification with a transmitted-light microscope.

J.E. Jankowski

During this process, exposed pollen grains on the microscope slides will be stained with
Safranin in order to highlight the morphological characteristics useful for their taxonomic
identification. Using a transmitted-light microscope, all grains present in each sample will be
counted. Identification using a scanning electron microscope (SEM) may be necessary for more
cryptic samples. Pollen taxa will be identified to genus or species using available literature on
the subject (e.g., Roubik and Moreno 1991, Herrera and Urrego 1996, Colinvaux et al. 1999,
Velasquez 1999) and the database developed by the Neotropical Paleoecology Research
Group (Bush and Weng 2007), as well as the modern pollen reference collection of the
Paleoecology Laboratory of Florida Institute of Technology.

Bird and flowering plant censuses-Data on range limitations in hummingbirds and
flowering plants come from censuses that have been conducted along the altitudinal gradient
from 2005-2008. For birds, I conducted point counts along the altitudinal gradient during the
breeding seasons (August November) of 2006-2008. In total 225 points were established
along trails and transects, averaging 17 points per 200m elevation. Point-count locations were
separated by 130m (horizontal distance) and marked with a GPS. Each count lasted ten
minutes, during which all birds detected by sight or sound were identified, and their distances
from the point were recorded. Counts at each point were repeated four times throughout the
breeding season. Point-count data will be combined with mist-netting survey data from the
gradient to determine the altitudinal ranges of the hummingbird species sampled. Understory
plants were sampled in 2007-2008 across 176 vegetation plots, each measuring 10x10m,
placed at each bird point-count location, in which every species >1m high was sampled.
Samples are stored and awaiting identification by Peruvian botanical experts in the lab of Dr.
Norma Salinas at the Universidad de San Antonio Abad del Cusco (UNSAAC). Data on tree
species distributions along the gradient were collected by Peruvian botanists within the labs of
collaborators Dr. Norma Salinas and of Dr. Miles Silman at Wake Forest University. A total of
15 1-hectare plots and 12 0.1-hectare plots were established at 250m intervals along the
altitudinal gradient. In each plot every stem l10cm dbh was identified and counted, and for
subplots within each plot, all understory stems >1m were counted and identified (or if field
identification was not possible for trees or understory stems, samples were collected and later
identified).

Data analysis-Dr. Mark Bush will provide a list of flowering-plant species from the pollen
grains identified in each sample for each of the 418 pollen samples collected. The extent to
which these samples represent the entire breadth of flowering plants pollinated will then be
determined for each species using saturation curves between number of samples and number

J.E. Jankowski

of additional flowering-plant species detected. We will find many pollen grains per pollen
sample, potentially representing many different species of flowering-plants. Seventeen of the
captured hummingbird species have eight or more pollen samples, and six species have at least
20 samples, which should allow us to identify a significant proportion of the flowering plants that
make up their diets. Five species have 30-50 pollen samples, allowing us to examine changes
in the flower use at different elevations within species' ranges. The pollen samples collected
are broadly distributed across the altitudinal range for each species, limiting the bias of any
single netting site in determining the list of flowering-plant species utilized.
The first hypothesis, that range limits in hummingbirds that are specialized to few flowering-
plant species coincide with limits in the distribution of their flowering plants, will be performed in
a two step process: (1) a preliminary visual examination by overlaying altitudinal distributions for
hummingbirds and their flowering-plants, followed by (2) a logistic regression to predict the
probability of occurrence of the hummingbird species along the altitudinal gradient, using the
occurrence of flowering-plant species as binary predictor variables (Agresti 1996).
Determining whether hummingbirds vary their use of flowering-plant species at different
altitudes will be analyzed by first developing a flowering-plant species list for each hummingbird
for different altitudinal zones (e.g., for each 200m zone). A similarity index (e.g., Sorenson's or
Morisita-Horn index) will be used to quantify the change in flowering-plant species composition
for each hummingbird across these altitudinal zones using the program EstimateS (Colwell
2005). This program offers indices that correct for undersampling biases (and allow sampling
biases to be assessed) and standard error estimates for the index, allowing for statistical
comparison of similarity values between different altitudinal zones.
The final research question, whether hummingbirds that are specialized on fewer species of
flowering-plant species have narrower altitudinal distributions, will be analyzed using a general
linear model (GLM) using the program SAS (SAS Institute, 2008). This analysis will be
performed for (1) all hummingbird species, (2) species with pollen sample sizes of eight or
more, and (3) species with pollen sample sizes of 17 or more. Within these groups, rarefaction
sub-sampling will be used to compare hummingbird species that differ in individual pollen
sample sizes.

Broader Impacts

Tropical mountains have been targeted as regions where climate change will have high
impacts on flora and fauna, forcing species to shift their ranges upslope (e.g., Pounds et al.
1999, Williams et al. 2003, Deutsch et al. 2008, Tewksbury et al. 2008). For conservation
initiatives, it is critical to describe the distribution and abundance of species in diverse montane
regions to generate a baseline of current species distributions and also identify potential factors
that influence species altitudinal range limits. Studies that project how species will shift their
ranges in response to different temperature regimes should consider species that may be
restricted by not only their allowable physiological responses but by species interactions like
competition or specialization to resources. This might be particularly relevant to hummingbirds
because of their often strong specializations with flowering plants and because most
hummingbird species live in tropical montane areas. Along this altitudinal gradient, for example,
86% of 64 total species have altitudinal range limits above 1000m (9 species have distributions
confined to elevations below 1000m, and only 4 species are exclusive to lowland elevations <
500m; Patterson et al. 2006). This study will provide a baseline for future studies to evaluate
how climate change affects altitudinal distributions in hummingbirds and the importance of
flowering-plant resources in driving range shifts in hummingbirds.
This proposed work represents an ongoing collaboration with the Neotropical
Paleoecology Laboratory, directed by Dr. Mark Bush at the Florida Institute of Technology,
whose focus has been to develop a reference collection and an online pollen database for

J.E. Jankowski

Neotropical flowering plants, with a strong concentration on Andean flora of southeastern Peru.
This collaboration has already fostered field research experience for an undergraduate student,
Zachariah Peterson, at Florida Institute of Technology, who worked with me during the
collection of the pollen samples in Manu Biosphere Reserve in 2006. Dr. Bush and I will create
another undergraduate research position in his lab to process the pollen samples. One of my
undergraduate assistants, Marcela Salazar, a Colombian student at the University of Florida, is
currently working on her honor's thesis with data from my dissertation research under my
mentorship, and will be a likely candidate for this position as she builds her research
experiences in preparation for graduate school. The pollen identification will be conducted by
Bryan Valencia, a trained palynologist from Cuzco, Peru. Any pollen samples that are new to
this Neotropical pollen collection will contribute to the lab's database. This is a great example of
international collaboration with scientists from the United States providing training and skill
development for students at both the beginning and later stages of their research careers.
A product of this research that will be available on the web is a hummingbird-flowering
plant photo guide, featuring the hummingbird species in this study for which pollen is being
analyzed, along with their primary flowering plants. This research is the result of a multi-lab
collaboration with botanists from the lab of Dr. Miles Silman in Wake Forest University and
paleoecologists in the Neotropical Paleoecology Lab. Zachariah Peterson has already begun
compiling bird and flowering plant photos that were taken during field work with me in Manu in
2006. This hummingbird-flowering plant guide will be especially useful for Peruvian and other
Latin American students who are beginning research projects in the Manu Biosphere Reserve,
especially for those who are interested in plant-pollinator interactions. Another product resulting
from this project will be a set of three compact discs of recorded bird species from the montane
forests of Manu Biosphere Reserve, representing foothill, middle- and high-elevation
communities (with 99 species per disc). These will be available as a learning resource for
researchers interested in studying birds of southeastern Peru. At the University of Florida,
songs are currently being organized and edited by an undergraduate student volunteering in our
lab.
Throughout my dissertation work, I have been committed to mentoring students from
Latin American and from underrepresented groups. During the prior field component of this
project (2005-2008), I have trained a variety of biology students with a range of ethnic
backgrounds, including eight female Peruvian biology students, two female Colombian
undergraduates, as well as two undergraduate students and one high school student from the
United States. Field skills acquired in this project include learning to identify tropical birds by
sight and sound, mist-netting and measuring birds, recording bird songs using digital equipment,
and censusing vegetation plots. Three of the Peruvian students, Rosalbina Butr6n Loayza,
Natividad Raurau Quisiyupanqui, and Maura Jurado Zevallos, are now completing their
"licenciatura" projects using data they collected under my mentorship. Rosalbina is determining
whether montane birds inhabiting bamboo patches (Guadua sp.), ranging from the Manu
foothills to lower cloud forest, are specialized to those habitats; until now, specialization to
bamboo communities has been considered a lowland phenomenon (Kratter 1997). Natividad is
identifying seeds from fecal samples of birds to determine the range of fruiting trees utilized by
different birds along the altitudinal gradient. Maura is planning a study for April 2009 to census
a high-elevation bird community during the non-breeding season. She will compare her data
with our breeding season dataset of species composition of the same location to determine
whether there is temporal variation in this bird community. Finally Marcela, the Colombian
undergraduate student mentioned above, is examining the potential for song mimicry between
closely related species that show species replacements along altitudinal gradients, as one
element of the interspecific territoriality that these species exhibit at their range boundaries.
Within my study region, I have also worked with the Asociaci6n para la Conservaci6n de
la Cuenca Amazonica (ACCA) to promote research at the Wayqecha Centro de Investigaci6n

J.E. Jankowski

(one of the primary field sites in my dissertation data collection). This has involved numerous
presentations to Peruvian university students from the Universidad de San Antonio Abad del
Cusco (UNSAAC), many of whom are aspiring tropical biologists. I have also presented my
work at science workshops for middle and high school children from the nearby towns of Patria
and Pilcopata. These presentations are of course in Spanish. I take pride in introducing
children to biology, birds, and the enormous species diversity that can be found in their
backyards. One of the most gratifying moments was when a young girl who told me that now
she wanted to become a biologist when she grew up.
I also take pride in being a role model for Latin American women students, who are
beginning the process of scientific investigation in a field where successful women tropical
biologists are in short supply. As a woman, I am part of an underrepresented group in the field
of tropical biology. I can think of only a couple prominent women biologists who study
Neotropical vertebrates and undertake extensive field seasons in remote locations. It has been
a challenge, yet very rewarding, to assemble the multi-cultural and multi-national research
teams necessary to conduct my dissertation research. The field work alone, which requires
months of living in tents and managing food and supplies into remote corners of montane forest,
has been a life-changing experience for all involved. Being able to accomplish this in a
Peruvian culture that typically looks to males for leadership and often looks down on women's
abilities has been an inspiration to women investigators of our research team, both national and
international.

o Undergraduate training in research (2005-present): I have mentored undergraduate
students in the field and lab, training them in methods of data collection and analysis,
teaching them how to use the scientific method, and helping them work on independent
research projects that they develop, mostly with topics related to my dissertation. Eight
of these students are female Peruvians, two are female Colombians (one with U.S.
citizenship), and four are undergraduate students from the United States (3 female, 1
male).

o Undergraduate teaching (2006-present): At the University of Florida, I have been a
teaching assistant for undergraduate courses, including Avian Biology (multiple
semesters; mostly upperclassmen) and Introductory Biology Laboratory (mostly
underclassmen). In my teaching, I reach students with a broad range of ethnic
backgrounds.

o Workshop leader with ACCA (2006-2007): With the Asociaci6n para la Conservaci6n
de la Cuenca Amazonica (ACCA) in Cusco, Peru, I led workshops in science and
environmental awareness for children in nearby towns, ages 8 to 14, many of whom
have never met or heard of a biologist before. This gave me the unique opportunity to
connect children to the nature in their backyards.

o Introducing high school students to scientific study (2007): I have worked with two
high school students in the lab, introducing them to scientific investigation and biology,
specifically working with microscopes and ectoparasite samples collected from birds
during my dissertation work. I brought another high school student to my field site in
southeastern Peru and trained him in data collection methods and conducting scientific
investigation.

Synergistic Activities
Head, Department of Ecology, Ethology, and Evolution at the University of Illinois: During my
administration we hired 3 women and 2 Hispanics to faculty positions (out of 6 positions), all of whom are
thriving.

Undergraduate student training: I typically employ 2-10 undergraduates each summer on research
projects relating to biodiversity. Three of my undergraduate advises (Judit Ungvari-Martin, Monique
Hiersoux, and Jane Bauer) are currently preparing manuscripts for eventual publication.

International Graduate student training: I have trained several international students, including
Muhammed Hussin (MS student from Malaysia, currently a professor in Malaysia), Miguel A. Marini
(Ph.D., currently a professor in the Federal University of Brasilia), Leonardo Chapa (Ph.D., currently a
professor in San Luis Potosi, Mexico), Juan Bouzat (Ph.D., an Argentinian currently an Assistant
Professor at Bowling Green), and Gustavo Londono (a Colombian co-advised with Doug Levey). I
currently have an Hispanic American graduate student, Ari Martinez, who works in Bolivia.

I work extensively with various federal (USFS,USFWS, NPS), state(lllinois DNR, Florida FWC), local
(Audubon Societies, NRDC), and private agencies(TNC, Environmental Defense) and with the media to
promote the conservation of neotropical migratory birds, principally by advising on land management and
acquisition.

Speaking on bird conservation at local public schools, Audubon Societies, and other organizations

Travel expenses:
I am requesting funds for traveling expenses for three visits to the Neotropical
Paleoecology Lab in Melbourne, Florida, to review the progress and results of the pollen
identification and consult with Dr. Bush. Melbourne is approximately 180 miles (360
miles round trip) from the University of Florida in Gainesville. Using University of
Florida's standard gas mileage costs of $0.44 per mile, and a two-day car rental for
approximately $64.20 (subcompact car from Avis with University employee discount)
travel expenses would yield $222.60 round trip for a single visit, and $667.80 for all three
trips.

Other Direct Costs:
Pollen identification: The primary expense in the proposed project is the
identification of pollen samples collected from hummingbirds. The pollen grains in each
sample will be identified in the Neotropical Paleoecology Laboratory of Dr. Mark Bush at
the Florida Institute of Technology. The costs for pollen analysis include processing
expenses for preparing the samples (e.g., chemical solutions used for remounting and
staining pollen grains) and lab consulting fees for the identification of the pollen grains.
Based on an estimated average of 1-2 hours to process and identify each sample, and
the amount of chemicals required, we have estimated a per sample expense of $9 for
processing and $15 for identification (also see letter of collaboration from Dr. Bush). For
418 samples, the total cost is $10,032.
Plant sample identification: Half of the plant samples that will be used to
determine flowering-plant species altitudinal ranges along the gradient are being stored
in the laboratory of Dr. Norma Salinas at the Universidad de San Antonio Abad de
Cusco, and require botanical experts within the lab to identify them. This consulting cost
for the botanical identification requires $10/day, and with plants from approximately 88
vegetation plots to identify, at an estimated rate of 1.5 plots per day, the total cost would
yield $586.67

The total cost of this project comes to $11,286.47.

Current and Pending Support
(See GPG Section II.C.2.h for guidance on information to include on this form.)
The following information should be provided for each investigator and other senior personnel Failure to provide this information may delay consideration of this proposal
Other agencies (including NSF) to which this proposal has been/will be submitted.
Investigator: Douglas Levey
Support: 0 Current H Pending 0 Submission Planned in Near Future D *Transfer of Support
Project/Proposal Title: International DDIG: What limits recruitment of tropical
trees? An experimental test of seed and establishment
limitation.

Support: 0 Current 0 Pending 0 Submission Planned in Near Future 0 *Transfer of Support
Project/Proposal Title: Catalyzing institutional change in STEM education at the
University of Florida

Source of Support: NSF
Total Award Amount: $ 1,000,000 Total Award Period Covered: 01/01/09 12/31/14
Location of Project: Gainesville, FL
Person-Months Per Year Committed to the Project. Cal:0.00 Acad: 0.00 Summ: 1.00
*If this project has previously been funded by another agency, please list and furnish information for immediately preceding funding period.

USE ADDITIONAL SHEETS AS NECESSARY

Page G-1

Current and Pending Support
(See GPG Section II.C.2.h for guidance on information to include on this form.)
The following information should be provided for each investigator and other senior personnel Failure to provide this information may delay consideration of this proposal
Other agencies (including NSF) to which this proposal has been/will be submitted.
Investigator: Douglas Levey
Support: 0 Current H Pending 0 Submission Planned in Near Future D *Transfer of Support
Project/Proposal Title: DISSERTATION RESEARCH: Using hummingbirds to test the link
between bird and plant distributions in tropical montane
forest

Source of Support:
Total Award Amount: $ Total Award Period Covered:
Location of Project:
Person-Months Per Year Committed to the Project. Cal: Acad: Summ:
*If this project has previously been funded by another agency, please list and furnish information for immediately preceding funding period.

Page G-2

USE ADDITIONAL SHEETS AS NECESSARY

Current and Pending Support
(See GPG Section II.C.2.h for guidance on information to include on this form.)
The following information should be provided for each investigator and other senior personnel Failure to provide this information may delay consideration of this proposal
Other agencies (including NSF) to which this proposal has been/will be submitted.
Investigator: Jill Jankowski
Support: 0 Current H Pending 0 Submission Planned in Near Future D *Transfer of Support
Project/Proposal Title: DISSERTATION RESEARCH: Using hummingbirds to test the link
between bird and plant distributions in tropical montane
forest

Source of Support:
Total Award Amount: $ Total Award Period Covered:
Location of Project:
Person-Months Per Year Committed to the Project. Cal: Acad: Summ:
*If this project has previously been funded by another agency, please list and furnish information for immediately preceding funding period.

Page G-3

USE ADDITIONAL SHEETS AS NECESSARY

Current and Pending Support
(See GPG Section II.C.2.h for guidance on information to include on this form.)
The following information should be provided for each investigator and other senior personnel Failure to provide this information may delay consideration of this proposal
Other agencies (including NSF) to which this proposal has been/will be submitted.
Investigator: Scott Robinson
Support: 0 Current H Pending 0 Submission Planned in Near Future D *Transfer of Support
Project/Proposal Title: DISSERTATION RESEARCH: Using hummingbirds to test the link
between bird and plant distributions in tropical montane
forest

Source of Support:
Total Award Amount: $ Total Award Period Covered:
Location of Project:
Person-Months Per Year Committed to the Project. Cal: Acad: Summ:
*If this project has previously been funded by another agency, please list and furnish information for immediately preceding funding period.

Page G-4

USE ADDITIONAL SHEETS AS NECESSARY

FACILITIES, EQUIPMENT & OTHER RESOURCES

FACILITIES: Identify the facilities to be used at each performance site listed and, as appropriate, indicate their capacities, pertinent
capabilities, relative proximity, and extent of availability to the project. Use "Other" to describe the facilities at any other performance
sites listed and at sites for field studies. USE additional pages as necessary.

Laboratory:

Florida Institute of Technology, Neotropical Paleoecology Lab, equipped
with Scanning Electron Microscope (SEM) and other equipment used for
pollen identification, located approximately 180 miles from the University
of Florida (see letter of collaboration from Dr. Mark Bush).

Clinical:

Animal:

Computer:

Office:

Other:

MAJOR EQUIPMENT: List the most important items available for this project and, as appropriate identifying the location and pertinent
capabilities of each.

Transmitted light microscope
Scanning electron microscope (SEM)

These items are located in the Neotropical Paleoecology Lab at Florida
Institute of Technology, directed by Dr. Mark Bush, a specialist in pollen
identification.

OTHER RESOURCES: Provide any information describing the other resources available for the project. Identify support services
such as consultant, secretarial, machine shop, and electronics shop, and the extent to which they will be available for the project.
Include an explanation of any consortium/contractual arrangements with other organizations.

FACILITIES, EQUIPMENT & OTHER RESOURCES

Continuation Page:

LABORATORY FACILITIES (continued):

NSF FORM 1363 (10/99)

Department of Biological Sciences
5rOjj1th ANNIVERSARY

Jill Jankowski
University of Florida Nov. 13th, 2008

Dear Jill,
Your project is an exciting one and we would be happy to continue our
collaboration. We have a photographic database of the flowers that the hummingbirds
were seen to visit during the 6 weeks that Zach Peterson spent in the field with you, and
also the associated pollen types. While this will not be a complete resource it should help
with the accuracy of some pollen identifications.
We should be able to provide identifications of pollen from the samples you have
collected. I will hire an undergraduate to do the processing and so an additional benefit of
your project will be an undergrad at Florida Tech being involved in science. As has been
our habit, we especially encourage women and minorities to apply for these positions.
The actual pollen identification will be done by one of my graduate students, probably by
Bryan Valencia (MS Florida Tech 2006) who is a native of Cuzco, Peru, and has 4 years
experience of Andean palynology.
The cost per sample will be $9 for processing and $15 for identification. While
some samples will be barren and very quick to do, others will require both light
microscopy and SEM to provide an optimum identification, hence this will probably
average out at 1-2 hours per sample. We generally have a fixed overhead rate of 15% for
this kind of consulting, but this can be waived as DDIG grants cannot carry overhead.
We would provide you with a sample-by-sample analysis of what we find, and
photographs of the pollen types. I believe that we could deal with all 300-500 samples
during the course of next summer.

My dissertation focuses on understanding patterns of species distributions in bird
communities along a tropical altitudinal gradient and the factors that maintain altitudinal range
limits in groups of species. I have conducted this investigation along an altitudinal gradient in
the Manu Biosphere Reserve of southeastern Peru because of this region's high species
diversity and the many possible distributional associations across species that could be
examined in detail along this gradient. In particular, I am addressing two distinctive patterns
found in birds in other tropical montane communities: (1) species replacements, or adjacent
non-overlapping altitudinal distributions, between closely related species, presumably
maintained by competitive interactions, which I tested experimentally using behavioral
responses to song playbacks; and (2) relationships between bird and plant communities, where
transitional zones in plant communities along the gradient (e.g., ecotones) might be reflected in
associated range limits in bird species' distributions as a response to sudden changes in
vegetation. The community-wide correspondence between birds and plants is being tested
using avian and plant census data collected from corresponding sites along the gradient.
Because it is difficult to examine mechanisms that limit species distributions when
considering all species of birds and plants, I have decided to focus on a subset of these
communities for which we might expect strong associations along gradients: hummingbirds and
flowering plants. If any group of birds were to show correspondence in their distributions and
range limits with plants along gradients, it would be this group. The proposed research takes a
novel approach that overcomes previous difficulties in examining resource specialization in
hummingbirds. Prior studies have used observations of hummingbirds at flowering plants in
their environments to describe plant-pollinator relationships. This approach is problematic,
especially in diverse tropical systems, because it requires repeated observations of all species
of flowering plants in a region. There is a simple solution-because hummingbirds carry, on
their bills and foreheads, pollen of the flowering plants they visit, one can sample and identify
the pollen from captured individuals to determine which flowers are being visited. With this in
mind, I collected pollen samples from the bills of captured hummingbirds during mist-netting
surveys along the altitudinal gradient. By collaborating with specialists that can identify the
pollen grains collected from each individual, I can create a list of flowering-plant species used by
each species of hummingbird. I will combine the resulting data detailing each hummingbird
species' diet with data from the bird and plant censuses already conducted to determine
whether hummingbird species' distributions and range limits coincide with those of their
flowering plants. Funds from the doctoral dissertation improvement grant, if awarded, would
allow the pollen identification to be completed, as the pollen analysis would otherwise be cost
prohibitive.

The research programs of my doctoral advisors overlap very little with the proposed research
project. While both of my advisors planted their roots in tropical ecology, neither has examined
the ecology of species range limits. Dr. Doug Levey's research fits under an umbrella of
evolutionary ecology of plant animal interactions and currently focuses on the effectiveness of
habitat corridors in conservation and impacts on the spatial ecology of animals, functions of
secondary compounds in ripe fruits, the importance of fruiting plants to birds, and the
evolutionary ecology of avian migration. Dr. Scott Robinson's research uses theoretical insights
of ecology and behavior to address questions of conservation significance in ecosystems of
North America and in the tropics. His current research projects address effects of forest and
grassland conservation on birds in the American Midwest, effects of urbanization on bird
communities, avian brood parasitism, and community organization in tropical forest birds of
white sands forest in the Amazon. Neither of my advisors is interested in seeking funding for the
required pollen analysis for the proposed research.